NeuroImage 16, 61–75 (2002)
doi:10.1006/nimg.2001.1046, available online at http://www.idealibrary.com on
A Developmental fMRI Study of the Stroop Color-Word Task
Nancy E. Adleman,* Vinod Menon,* ,† ,‡ Christine M. Blasey,* Christopher D. White,*
Ilana S. Warsofsky,* Gary H. Glover,§ and Allan L. Reiss* ,†
*Departments of Psychiatry & Behavioral Sciences, †Program in Neuroscience, ‡Stanford Brain Research Center,
and §Radiology, Stanford University School of Medicine, Stanford, California 94305
Received April 30, 2001
We used fMRI to investigate developmental changes
in brain activation during a Stroop color-word inter-
ference task. A positive correlation was observed be-
tween age and Stroop-related activation (n ⴝ 30) in the
left lateral prefrontal cortex, the left anterior cingu-
late, and the left parietal and parieto-occipital corti-
ces. No regions showed a negative correlation between
activation and age. We further investigated age-re-
lated differences by stratifying the sample into three
age groups: children (ages 7–11), adolescents (ages 12–
16), and young adults (ages 18 –22). Young adult sub-
jects (n ⴝ 11) displayed significant activation in the
inferior and middle frontal gyri bilaterally, the left
anterior cingulate, and bilateral inferior and superior
parietal lobules. Between-group comparisons re-
vealed that young adults had significantly greater ac-
tivation than adolescent subjects (n ⴝ 11) in the left
middle frontal gyrus and that young adults showed
significantly greater activation than children (n ⴝ 8)
in the anterior cingulate and left parietal and parieto-
occipital regions, as well as in the left middle frontal
gyrus. Compared to children, both adult and adoles-
cent subjects exhibited significantly greater activa-
tion in the parietal cortex. Adult and adolescent
groups, however, did not differ in activation for this
region. Together, these data suggest that Stroop task-
related functional development of the parietal lobe
occurs by adolescence. In contrast, prefrontal cortex
function contributing to the Stroop interference task
continues to develop into adulthood. This neuromatu-
rational process may depend on increased ability to
recruit focal neural resources with age. Findings from
this study, the first developmental fMRI investigation
of the Stroop interference task, provide a template
with which normal development and neurodevelop-
mental disorders of prefrontal cortex function can be
assessed. © 2002 Elsevier Science (USA)
INTRODUCTION
Since its introduction in 1935, the Stroop Color-Word
task has been a classic measure of frontal lobe function
61
(MacLeod, 1991). In this task, the subject is instructed
to name the color of the ink of an incongruent word-
color stimulus (i.e., RED printed in green ink). Because
word reading is a more automatic cognitive process
than color naming, subjects must resolve cognitive in-
terference and inhibit the incorrect, but more facile,
response in order to properly respond. This response
competition commonly results in an increase in re-
sponse time (RT) on incongruent trials as compared to
congruent word-color pairs (i.e., RED printed in red
ink) or neutral pairs (i.e., DOG printed in red ink, or
XXX printed in red ink).
The cognitive processes underlying the Stroop task
(i.e., response inhibition, interference resolution, and
behavioral conflict resolution) are considered executive
processes mediated by the frontal lobe. Patients with
lateral prefrontal lobe lesions commit more errors on
the Stroop task than do normal controls (Vendrell et
al., 1995). Perret (1974) proposed a left hemisphere
frontal location for interference resolution in the
Stroop task after studying patients with frontal lobe
lesions. Because of its critical dependence on frontal
lobe function, the Stroop task is an ideal tool for study-
ing typical and atypical development of executive pro-
cesses. Several studies have utilized the Stroop task
(Dash and Dash, 1982) and modified Stroop tasks (Di-
amond and Taylor, 1996; Gerstadt et al., 1994) to ex-
plore the maturation of executive processes, and have
found that such processes develop with age.
In conjunction with the behavioral development of
executive function, researchers have recently provided
evidence for postadolescent maturation of the frontal
lobes (Sowell et al., 1999). Giedd et al. (1999) found that
frontal gray matter increases during preadolescence,
peaks during adolescence, and decreases during post-
adolescence. Although there has been evidence of
structural frontal lobe maturation as well as behav-
ioral maturation of executive processes during child-
hood, no functional imaging studies to date have used
the Stroop task to investigate the relationship between
frontal lobe function and cognition from a developmen-
tal standpoint.
1053-8119/02 $35.00
© 2002 Elsevier Science (USA)
All rights reserved.
62 ADLEMAN ET AL.
Although there have been no developmental imaging
studies of the Stroop task to date, data from several
Stroop task imaging studies have been reported in
adults (Bench et al., 1993; Carter et al., 1995; Larrue et
al., 1994; Pardo et al., 1990; Peterson et al., 1999;
Taylor et al., 1997). However, considerable variability
exists in the brain regions observed to be activated by
the Stroop task. For example, utilizing a similar non-
word control task as that used in the present study,
Bench et al. (1993) reported Stroop task-related acti-
vation in right orbitofrontal cortex, right frontal polar
cortex, and right anterior cingulate. Taylor et al.
(1997), also employing a nonlexical control task, found
activation in the right middle frontal gyrus (MFG), left
inferior frontal gyrus (IFG), left parietal region, and
left insula; no activation was observed in the anterior
cingulate cortex (ACC). In summary, most investiga-
tions of the Stroop task have found activation in at
least one of three main regions: the ACC (Carter et al.,
1995; Larrue et al., 1994; Pardo et al., 1990; Peterson et
al., 1999), parietal lobe (Bench et al., 1993; Carter et
al., 1995; Peterson et al., 1999; Taylor et al., 1997), and
lateral prefrontal cortex (Bench et al., 1993; Carter et
al., 1995; Pardo et al., 1990; Taylor et al., 1997). The
goal of the present study was to employ the classic
Stroop interference task with functional magnetic res-
onance imaging (fMRI) to investigate neuromatura-
tional processes underlying cognitive development in
individuals ranging in age from early childhood to
young adulthood. In accord with past Stroop task neu-
roimaging research, we hypothesized that develop-
ment of executive processes involved in the Stroop
interference process would be localized to these three
regions.
In contrast to most functional imaging studies of
cognitive development, which typically employ only
two groups of subjects, children and adults (e.g.,
Thomas et al. (1999)), the present study utilized three
developmental age groups (children, adolescents, and
young adults), thus permitting a more fine-grained
investigation of the trajectory of cognitive developmen-
tal changes. In particular, the inclusion of an adoles-
cent group is a potentially significant addition as it is
well known that the onset of puberty affects both de-
velopment (Giedd et al., 1999) and behavior (Buchanan
et al., 1992).
This study is a component of our laboratory’s ongoing
efforts to provide an initial template of neurofunctional
maturation underlying key components of cognitive
development during childhood. Broadening our knowl-
edge base pertaining to the trajectory and variability of
typical brain development in children will eventually
permit a more precise interpretation of individual
brain development and function in children with atyp-
ical cognitive development.
MATERIALS AND METHODS
Subjects
Subjects were recruited from the greater San Fran-
cisco Bay area. Two exclusion criteria were applied
when screening for potential participants: (1) evidence
of identifiable cognitive impairment or (2) the presence
of a behavioral/psychological disorder. Cognitive func-
tioning was assessed using the WISC-III (ages 7
through 16) or WAIS-III (ages 16 through 22). Persons
who scored lower than 80 for full scale IQ were ex-
cluded from further participation. The presence of be-
havioral/psychological impairments was assessed us-
ing either the Symptom Checklist-90-R (SCL-90-R)
(ages 13 through 22) (Derogatis and Savitz, 1999) or
the Child Behavioral Checklist (CBCL) (ages 7 through
18) (Achenbach, 1991). Subjects with significant eleva-
tions (1 SD ⬎ population mean) on these instruments
were excluded from the study.
Thirty healthy, right-handed, biologically indepen-
dent subjects met these selection criteria and partici-
pated in the Stroop task fMRI study after giving writ-
ten informed consent. The total sample included 10
males and 20 females ranging in age from 7 to 22 years
(mean age: 15.42, SD ⫽ 4.24). Because of behavioral
literature that suggests that by the age of seven chil-
dren have developed their reading skills to the point
that they experience semantic interference (Rosinski,
1977; Rosinski et al., 1975) and that children under the
age of seven do not show interference effects (Comalli
et al., 1962), we confined our subject pool to children
aged seven and older.
Subjects were divided into three age groups with
similar male to female proportions. The group of chil-
dren consisted of 3 males and 5 females (n ⫽ 8) ranging
in age from 7.66 to 11.96 years (mean ⫽ 10.16, SD ⫽
1.66). The adolescent group included 4 males and 7
females (n ⫽ 11) ranging from 12.58 to 16.85 years of
age (mean ⫽ 14.68, SD ⫽ 1.27). The young adult group
consisted of 3 male and 8 female subjects (n ⫽ 11),
ranging in age from 17.39 to 22.68 years (mean ⫽
19.98, SD ⫽ 1.72).
Neuropsychological Stroop Test
Because of head movement that would result from
fully vocalizing responses within the scanner, we used
behavioral data acquired outside the scanner to inves-
tigate age-related changes in performance of the
Stroop task. In the present study, a standard Stroop
Color-Word task (Golden, 1978) was administered to
every subject. The Stroop test includes 3 time-limited
(45-s) subtests. First, subjects are asked to read a list of
words printed in black ink that name colors (i.e., “red,”
“green,” “blue”). Second, subjects are presented with a
list of “XXXXX”s that differ in ink color. Subjects name
the color of the ink for each “XXXXX.” In the third
 DEVELOPMENTAL fMRI OF THE STROOP TASK 63

TABLE 1
Results of Multiple Regression Analysis Examining the Relationship between FSIQ, Age, and Stroop Subscores

Zero-order Partial Part Beta

Dependent variable Predictor correlation correlation correlation R 2 delta Beta P value

Word raw score FSIQ 0.191 ⫺0.092 ⫺0.067 0.037 ⫺0.072 0.635
AGE 0.685 0.674 0.661 0.437 0.712 ⬍0.001
Color raw score FSIQ 0.340 0.059 0.032 0.116 0.035 0.761
AGE 0.837 0.814 0.765 0.585 0.824 ⬍0.001
Color–word raw score FSIQ 0.496 0.357 0.221 0.246 0.238 0.057
AGE 0.786 0.747 0.649 0.421 0.698 ⬍0.001
Interference raw score FSIQ 0.514 0.434 0.401 0.246 0.432 0.019
AGE 0.382 0.240 0.206 0.043 0.222 0.209

subtest, subjects are then presented with a list of the
words “red,” “green,” and “blue;” in this presentation,
however, the ink color of the words is discordant with
the presented word (i.e., the word “red” printed in blue
ink). Again, subjects are asked to say the color of the
ink and ignore the word’s semantic meaning (i.e., re-
spond “blue” instead of “red” in the previous example).
fMRI Stroop Task
The experiment presented in the scanner began with
a 30-s rest epoch followed by 3 cycles of alternating
30-s epochs of experimental trials and control trials, a
30-s rest epoch, another 3 cycles of control and exper-
imental epochs, and a final 30-s rest epoch. In each
epoch, there were 15 stimuli, each presented for 1350
ms, with a total interstimulus interval (ISI) of 2000 ms.
In order to highlight the complete network of brain
regions involved in Stroop task-related cognitive com-
ponents and their development, we employed a low-
level nonlinguistic baseline as the control condition
that only controlled for the process of color naming.
The control stimuli consisted of a string of 3, 4, or 5
colored “X”s, presented in red, blue, or green. The sub-
jects were asked to subvocalize the color of the “X”s

TABLE 2
Brain Areas That Showed Significant Activation in (1) Linear Correlation between Age and Activation; (2) Young Adult
Group Average; (3) Adult versus Child Comparison; (4) Adult versus Adolescent Comparison; (5) Adolescent versus Child
Comparison

No. of Cluster Z max Primary peak

Activated Region voxels P (cor) primary peak location

Linear correlation between age and activation

Left anterior cingulate cortex and lateral prefrontal regions 6109 ⬍0.001 4.57 ⫺4, 14, 28 L ACC (BA 24/32)
(SFG/MFG)
Left inferior parietal lobe, parieto-occipital regions, cuneus, 1700 ⬍0.001 3.67 ⫺26, ⫺62, 16 L PCC/cuneus
lingual gyrus, posterior cingulate cortex (BA31/7)
Young adult group average
Bilateral MFG, IFG, and anterior cingulate cortex 12030 ⬍0.001 4.37 ⫺34, 22, 34 L MFG (BA 8)
Right inferior parietal lobe, supramarginal gyrus, 1681 0.001 4.25 60, ⫺46, 28 R
superior/medial occipital gyrus IPL/supramarginal gyrus
(BA 40)
Left inferior and superior parietal lobe 3474 ⬍0.001 4.03 ⫺28, ⫺68, 52 L SPL (BA 7)
Adult–child
Anterior cingulate cortex 827 0.049 4.12 ⫺6, 26, 16 L ACC (BA 24)
Left MFG and SFG 2511 ⬍0.001 3.92 ⫺28, 14, 50 L MFG (BA 6)
Left parieto-occipital regions: lingual gyrus, precuneus, cuneus 937 0.025 3.58 ⫺14, ⫺66, ⫺2 L lingual gyrus
(BA 18/19)
Adult–adolescent
Left MFG, ACC, SFG 1376 0.002 3.81 ⫺28, 12, 52 L MFG (BA 6)
Adolescent–child
Left cuneus, posterior cingulate gyrus, precuneus 1858 ⬍0.001 4.11 ⫺14, ⫺74, 22 L cuneus (BA
17/18)

Note. For each cluster, the activated region(s), number of voxels activated, corrected P value, maximum Z score, and location of peak
activation are shown. ACC, anterior cingulate cortex; MFG, middle frontal gyrus; SPL, superior parietal lobe; IPL, inferior parietal lobe; PCC,
posterior cingulate cortex; IFG, inferior frontal gyrus; SFG, superior frontal gyrus.
64 ADLEMAN ET AL.

FIG. 1. Brain regions that showed significant (P ⬍ 0.05) Stroop task-related activation in the young adult group include the bilateral
frontal regions and bilateral parietal and parieto-occipital regions. Activated regions include the middle frontal gyrus, inferior frontal gyrus,
inferior parietal lobe, superior parietal lobe, and the superior/medial occipital gyrus. fMRI activation was superposed on the average of the
11 young adult subjects’ individual T1-weighted images normalized to Talairach space.

quietly to themselves. The experimental stimuli were
incongruent color-word pairs, in which the print of the
ink did not match the color specified by the word (i.e.,
the word “red” printed in green). The subjects were
asked to identify, and say quietly to themselves, the
color of the ink in which the word was printed, and not
to read the word itself (in our example, to say “green”).
All subjects were trained on the task before scanning
until they showed and expressed both understanding
and compliance.
fMRI Acquisition
Images were acquired on a 1.5T GE Signa scanner
with Echospeed gradients using a custom-built whole
head coil that provides a 50% advantage in signal to
 DEVELOPMENTAL fMRI OF THE STROOP TASK 65

FIG. 2. Brain areas in which Stroop task-related activation was significantly (P ⬍ 0.05) correlated with age. (A) fMRI activation was
superposed on the average of 29 individual T1-weighted images normalized to Talairach space. (B) Surface rendering of fMRI activation on
a T1-weighted Montreal Neurological Institute (MNI) template image.
66 ADLEMAN ET AL.
noise ratio over that of the standard GE coil (Hayes
and Mathias, 1996). A custom-built head holder was
used to prevent head movement. Eighteen axial slices
(6 mm thick, 1 mm skip) parallel to the anterior and
posterior commissure covering the whole brain were
imaged with a temporal resolution of 2 s using a T2*-
weighted gradient echo spiral pulse sequence (TR ⫽
2000 ms, TE ⫽ 40 ms, flip angle ⫽ 89° and 1 interleave)
(Glover and Lai, 1998). The field of view was 240 mm
and the effective inplane spatial resolution was 3.75
mm. To aid in localization of functional data, a high
resolution T1-weighted spoiled grass gradient recalled
(SPGR) 3-D MRI sequence with the following parame-
ters was used: TR ⫽ 24 ms; TE ⫽ 5 ms; flip angle ⫽ 40°;
124 slices in coronal plane; 256 ⫻ 192 matrix; acquired
resolution ⫽ 1.5 ⫻ 0.9 ⫻ 1.2 mm. The images were
reconstructed as a 124 ⫻ 256 ⫻ 256 matrix with a 1.5 ⫻
0.9 ⫻ 0.9 mm spatial resolution. Structural image av-
erages were made for the entire subject pool as well as
for each age group. Because of excessive movement
during one structural scan (an individual in the child
group), only 29 subjects were included in the average
structural image.
The task was programmed using Psyscope (Cohen et
al., 1993) on a Macintosh (Sunnyvale, CA) notebook
computer. Initiation of scan and task was synchronized
using a TTL pulse delivered to the scanner timing
microprocessor board from a “CMU Button Box” micro-
processor (http://poppy.psy.cmu.edu/psyscope) con-
nected to the Macintosh. Stimuli were presented visu-
ally at the center of a screen using a custom-built
magnet compatible projection system (Resonance
Technology, CA).
Image Preprocessing
Images were reconstructed, by inverse Fourier
transform, for each of the 225 time points into 64 ⫻
64 ⫻ 18 image matrices (voxel size: 3.75 ⫻ 3.75 ⫻ 7
mm). fMRI data were preprocessed using SPM99b
(http://www.fil.ion.bpmf.ac.uk/spm). Images were cor-
rected for movement using least square minimization
without higher-order corrections for spin history and
normalized to MNI coordinates. Images were then re-
sampled every 2 mm using sinc interpolation and
smoothed with a 4-mm Gaussian kernel to decrease
spatial noise. MNI coordinates were transformed to
Talairach (Tal) coordinates using a nonlinear transfor-
mation (Brett, 2000, http://www.mrc-cbu.cam.ac.uk/
Imaging/mnispace.html).
Statistical Analysis
Behavioral data. Raw scores on the Stroop task
were calculated by determining the number of items
completed within the 45-s time interval on each of the
three subtests (color, word, color–word). An interfer-
ence score was derived from the prediction of perfor-
mance on the third task (based on performance on the
first two tasks) minus the actual performance. Raw
interference was calculated using the same equation
the Stroop neuropsychological test provides for inter-
ference calculation. 1 For our experimental purposes,
this calculation of an interference score was derived
from the three subtest raw scores rather than from the
age-corrected scores provided in the Stroop manual
(which are described as “experimental” in the manual)
(Golden, 1978). The benefits of using raw scores in
cross-sectional studies of development have been dis-
cussed (Kraemer et al., 2000) and have been used in-
stead of age-standardized scores in similar investiga-
tions (Bettner et al., 1971).
Multiple regression analyses were performed to as-
sess the predictive value of age and full-scale IQ
(FSIQ), as measured by the WISC-III or WAIS-III, on
the Stroop performance measures.
fMRI data analysis. Statistical analysis was per-
formed on individual and group data using the general
linear model and the theory of Gaussian random fields
as implemented in SPM99b (Friston et al., 1995). This
method corrects for temporal and spatial autocorrela-
tions in the fMRI data (Worsley and Friston, 1995).
In the first step of analysis, a within-subject proce-
dure was used to model all the effects of interest. The
individual subject models were identical across sub-
jects (i.e., a balanced design was used). Confounding
effects of fluctuations in global mean were removed by
proportional scaling where, for each time point, each
voxel was scaled by the global mean at that time point.
Low frequency noise was removed with a high pass
filter (0.5 cycles/min) applied to the fMRI time series at
each voxel. A temporal smoothing function (Gaussian
kernel corresponding to dispersion of 8 s) was applied
to the fMRI time series to enhance the temporal signal
to noise ratio. The effects of interest for each subject
were then defined with the relevant contrasts of the
parameter estimates. For each of these contrasts, a
corresponding contrast image was made. Voxel-wise t
statistics were normalized to Z scores to provide a
statistical measure of activation independent of sam-
ple size. Finally, in order to determine the presence of
significant clusters of activation, the joint expected
probability distribution of height and extent of Z scores
(Poline et al., 1997), with height (Z ⬎ 1.67; P ⬍ 0.05)
and extent threshold (P ⬍ 0.05), was used in order to
correct for spatial correlations in the data.
Activation analysis. Group analysis was performed
using a random-effects model. The random-effects
model estimates the error variance for each condition
of interest across subjects, rather than across scans
1
The equations employed were as follows: (W ⴱ C)/(W ⫹ C) ⫽ CW⬘;
CW ⫺ CW⬘ ⫽ RAWINTF. W, raw word score; C, raw color score; CW,
raw color-word score.
 DEVELOPMENTAL fMRI OF THE STROOP TASK
(Holmes and Friston, 1998) and therefore provides a
stronger generalization to the population. This analy-
sis proceeded in two steps. In the first step, contrast
images for each subject and each effect of interest were
generated as described above. In the second step, these
contrast images were used to examine the correlation
between age and brain activation and between-group
activation differences as described below. We initially
analyzed data from young adult subjects to determine
the consistency of our findings with those previously
reported for adult samples.
Correlation between age and activation. Linear re-
gression was used to determine voxels that showed
either positive or negative age-related changes during
the Stroop interference task (interference vs color nam-
ing condition contrast) across all subjects. Subject age
was used as a covariate of interest and FSIQ was used
as a nuisance covariate due to unexpected group dif-
ferences for this measure. Voxel-wise t statistics were
computed for the covariate of interest. The t statistics
were then normalized to Z scores. Significant clusters
of activation were determined using the joint expected
probability distribution of height and extent of Z
scores, with height (Z ⬎ 1.67; P ⬍ 0.05) and extent
threshold (P ⬍ 0.05).
Group differences in Stroop task-related activation.
In order to further delineate the trajectory of develop-
mental changes, Stroop task-related activation differ-
ences between the three groups (children, adolescents,
and young adults) were compared. Individual contrast
images reflecting task-related activation were first cre-
ated for each subject. Groups were compared using
two-sample, two-tailed, t tests after covarying out dif-
ferences in full scale IQ. The following group compar-
isons were made: (1) young adult versus child; (2)
young adult versus adolescent; and (3) adolescent ver-
sus child.
RESULTS
IQ
Although age was not significantly correlated with
verbal IQ (r ⫽ 0.291; P ⫽ 0.118) or performance IQ (r ⫽
0.297; P ⫽ 0.111), linear regression indicated a signif-
icant correlation between age and full-scale IQ (r ⫽
0.37; P ⫽ 0.044). This significant age-effect in IQ was
related to a sampling bias in the young adult group due
to the inclusion of undergraduates at Stanford Univer-
sity. Because of these age-related findings, we used
full-scale IQ as a nuisance covariate in all analyses of
group and age-related effects.
Stroop Task Performance
Multiple regression analyses (Table 1) showed that
age explained a significant proportion of the variance
in Stroop task subtest scores (word raw: beta ⫽ 0.712,
67
P ⬍ 0.001; color raw: beta ⫽ 0.824, P ⬍ 0.001; color–
word raw: beta ⫽ 0.698, P ⬍ 0.001). However, age was
not significantly correlated with raw interference score
(P ⫽ 0.209). In contrast, FSIQ did not account for a
significant proportion of the variance in word raw (P ⫽
0.635), color raw (P ⫽ 0.761), or color–word raw scores
(P ⫽ 0.057), but was a significant predictor of raw
interference score (P ⫽ 0.019).
Brain Activation
Young adult group activation. Young adults
showed significant Stroop task-related activation bilat-
erally in the IFG, the MFG and the anterior cingulate
(Z max ⫽ 4.37, P ⬍ 0.001), bilateral inferior parietal
lobe/supramarginal gyrus (R side: Z max ⫽ 4.25, P ⫽
0.001; L side: Z max ⫽ 4.03, P ⬍ 0.001), and the left
superior parietal lobe (Z max ⫽ 4.03, P ⬍ 0.001; for
summary of all activation see Table 2 and Fig. 1).
Activation was also found to extend into the caudate
(Tal x, y, z ⫽ 16, 10, 6 mm in Fig. 1).
Correlation between age and brain activation. After
covarying for FSIQ, two clusters of brain activation
were significantly correlated with age (Table 2; Fig. 2).
The first cluster spanned the left anterior cingulate
gyrus (BA 24/32) and prefrontal regions including the
left superior frontal gyrus (SFG) (BA 6) and bilateral
MFG (BA 9) (Z max ⫽ 4.57, P ⬍ 0.001). Activation from
this cluster also was found to extend into bilateral
caudate/putamen (Tal x, y, z ⫽ 16, 10, 6 mm in Fig. 2A).
The second cluster spanned the left posterior cingulate
gyrus and cuneus (BA 31/7), the left lingual gyrus (BA
18/19), and the left inferior parietal lobe (BA 40) (Z
max ⫽ 3.67, P ⬍ 0.001).
In order to further explore the relationship between
age and focal activation in these two regions, a corre-
lational analysis (n ⫽ 30) was run between age and
activation level (arbitrary measure) in the foci maxima
of the two regions. The foci maxima were located in (1)
the left ACC (BA 24/32, Tal x, y, z ⫽ ⫺4, 14, 28 mm),
and (2) the left posterior cingulate cortex (PCC)/cuneus
(BA 31/7, Tal x, y, z ⫽ ⫺26, ⫺62, 16 mm). The activa-
tion level in both of these foci was significantly corre-
lated with subject age (P ⬍ 0.0001, see Figs. 3 and 4).
Young adults vs children. Young adults showed
greater Stroop task-related activation, as compared to
children, in the left and right anterior cingulate gyrus
(BA 24/32) (Z max ⫽ 4.12, P ⫽ 0.049), the left MFG and
SFG (BA 6/9/45) (Z max ⫽ 3.92, P ⬍ 0.001), and left
occipitoparietal regions, including the lingual gyrus
(BA 18/19), precuneus (BA 31/7), and the cuneus (BA
17) (Z max ⫽ 3.58, P ⫽ 0.025; for summary of all
activation see Table 2 and Fig. 5). Children did not
show significantly greater Stroop task-related activa-
tion than young adults in any brain region.
Young adults vs adolescents. Young adults had sig-
nificantly greater activation during the Stroop task, as
68 ADLEMAN ET AL.

FIG. 3. Correlation (n ⫽ 30) between age and activation level (arbitrary scale) in the peak voxel of activation in the left anterior cingulate
cortex (BA 24/32, Tal x, y, z ⫽ ⫺4, 14, 28 mm, P ⬍ 0.0001).
FIG. 4. Correlation (n ⫽ 30) between age and activation level (arbitrary scale) in the peak voxel of activation in the left posterior cingulate
cortex/cuneus (BA 31/7, Tal x, y, z ⫽ ⫺26, ⫺62, 16 mm, P ⬍ 0.0001).
 DEVELOPMENTAL fMRI OF THE STROOP TASK 69

FIG. 5. Brain areas showing significantly greater Stroop task-related activation in the young adults, compared to the children, included
the anterior cingulate bilaterally, the left MFG, the lingual gyrus, precuneus, and cuneus. Other details as described in the legend of Fig. 2.

compared to adolescents, in the left middle frontal gy-
rus (BA 6/8/9) (Z max ⫽ 3.81, P ⫽ 0.002; for summary
of all activation see Table 2, Fig. 6). Adolescents did not
show significantly greater Stroop task-related activa-
tion than young adults in any brain region.
Adolescents vs children. Adolescents exhibited sig-
nificantly greater activation than children in the left
cuneus (BA 17/18) and the posterior cingulate gyrus/
precuneus (BA 31) (Z max ⫽ 4.11, P ⬍ 0.001; for
summary of all activation see Table 2, Fig. 7). Children
showed greater activation than adolescents in two sig-
nificant clusters: the first included the right cerebel-
lum, right hippocampus, and midbrain; the second
cluster was localized in the right superior temporal
gyrus (STG) (BA 22/42).
DISCUSSION
Improved understanding of the dynamic and vari-
able nature of brain development and function
throughout the human lifespan is of the highest scien-
tific priority. In particular, elucidating typical develop-
70 ADLEMAN ET AL.
mental processes during childhood and adolescence
will enhance our ability to recognize and treat atypical
development and disease processes in the future.
The present study is the first to describe the rela-
tionship between Stroop task-related brain activation
and age, as well as the first to provide evidence for
ongoing developmental changes in Stroop-processing
brain networks that correspond to improvements in
behavioral task performance. A major strength of the
present study was the voxel-by-voxel correlation anal-
ysis using a random effects model (Holmes and Friston,
1998). We examined both positive and negative age-
related changes in brain activation during perfor-
mance of the Stroop task. This allowed us to examine
whether there is an increase in activation with age
corresponding to an ability to recruit appropriate re-
sources for task performance or a decrease in activa-
tion corresponding to better optimization and de-
creased task difficulty with age.
There are three major processes involved in perfor-
mance of the Stroop Color–Word task: word reading,
color naming, and interference resolution. In order to
fully characterize the development of brain activity
during Stroop task performance, we used a control task
involving color naming. Using this nonlexical control
task (colored “XXXXX”s) allowed us to examine the
development of interference resolution in the context of
word reading.
Activation in Young Adults
Young adults showed significant Stroop task-related
activation bilaterally in the IFG, MFG, inferior pari-
etal lobe/supramarginal gyrus, superior parietal lobe,
and the left anterior cingulate. This pattern of brain
activation in adults is consistent with Stroop task ac-
tivation revealed in other adult neuroimaging studies
(Pardo et al., 1990; Larrue et al., 1994; Carter et al.,
1995; Peterson et al., 1999; Bench et al., 1993). These
data, combined with those of the literature, provided a
template against which developmental-related differ-
ences in activation could be assessed in the two
younger age groups.
Linear Changes with Age
Both behavioral performance and patterns of brain
activation showed age-related changes. Raw word,
color, and color–word scores showed linear increases
with age, reflecting cognitive maturation in several
major processes associated with the Stroop task (read-
ing, color identification, and resolution of cognitive in-
terference). The behavioral results of this study are
consistent with previous behavioral studies that have
reported improvement in word reading skill with age.
Basic reading skills, as measured by the Woodcock-
Johnson III, are shown to markedly increase between
the ages of 5 and 25 years (McGrew and Woodcock,
2001). Analyses from the behavioral Stroop task in the
present study, also show a significant relationship be-
tween age and word reading skill, which develops over
the course of childhood and adolescence.
A positive correlation also was observed between age
and activation in the left lateral prefrontal cortex, the
left anterior cingulate, and the left parietal and parie-
to-occipital cortices. This linear correlation was most
significant in the left lateral prefrontal cortex and left
parieto-occipital regions. The left lateral prefrontal
cluster consisted of a much larger cluster of voxels (No.
of voxels ⫽ 6109) than the left parieto-occipital cluster
(No. of voxels ⫽ 1700). No regions showed decreased
activation with age. These results suggest that the
developmental trajectory of cognitive processing
needed for the Stroop task is characterized by increas-
ing ability to recruit additional neural resources.
Several recent developmental imaging studies of
frontal lobe function have shown increased, more dif-
fuse, activation in children as compared to adults (Ca-
sey et al., 1997; Gaillard et al., 2000; Thomas et al.,
1999). At first glance, these results may seem to con-
tradict our findings of increased Stroop task-related
activation with age. These differences may arise partly
from methodological differences. Previous studies have
used broadly defined regions of interest (ROIs); for
example, in a developmental study of the gonogo, Ca-
sey et al. (1997) examined activation in the entire ACC,
SFG, MFG, and IFG. In contrast, the present study
used a voxel-by-voxel regression analysis to examine
the precise voxels in which Stroop task-related activa-
tion changes with age. Thus, it is possible that there
are focal increases in activation that are not captured
using broadly defined ROIs. It is also possible that
there is more diffuse activation in children, which be-
comes more focal with age in very specific brain re-
gions.
Subjects showed increases in behavioral perfor-
mance in conjunction with increases in regional acti-
vation. The increase in activation with age may be
correlated with simultaneous improvements in behav-
ioral performance. Rubia et al. (2000) studied normal
adolescents and adults performing a motor timing task
and found that better delay-task performance in adults
paralleled increased activation in a brain network in-
cluding prefrontal and parietal cortical regions and the
putamen. Below, we further discuss age-related
changes in activation during performance of the Stroop
task by specific brain region (lateral prefrontal cortex,
parieto-occipital cortex, and anterior cingulate cortex).
We discuss these results with regard to both linear
age-related changes as well as differences between
stratified age groups (young adult, adolescent, and
child).
 DEVELOPMENTAL fMRI OF THE STROOP TASK
Lateral Prefrontal Cortex
Data from the present study provide evidence for
functional development in lateral prefrontal cortex re-
lated to the Stroop task. Prefrontal regions in which
activation was significantly correlated with age in-
cluded bilateral MFG (BA 9) and left SFG (BA 6). Left
MFG and SFG also showed significant differences in
activation in both the adult– child comparison, and the
adult–adolescent comparison, suggesting that the neu-
ral processes associated with prefrontal cortex activa-
tion develop throughout childhood, adolescence, and
into young adulthood.
Age-related differences in prefrontal activation were
specifically localized to the left hemisphere. Left hemi-
sphere differences in activation are consistent with the
verbal nature of the Stroop task. Due to the low-level
baseline control task, some of the age-related changes
in functional activation may reflect increasing word
reading proficiency. In the present study, no age-re-
lated activation changes were seen in the right prefron-
tal cortex. In particular, we did not see increased right-
sided activation in young children as compared to
adolescents and young adults.
Previous lesion and imaging studies have implicated
lateral prefrontal regions in both interference process-
ing/response inhibition (Casey et al., 1997; Pujol et al.,
2000) and word reading/production (Cabeza and Ny-
berg, 2000). Pujol et al. (2000), in a study of demyeli-
nating lesions in patients with multiple sclerosis, re-
ported that 45% of the variance in Stroop interference
time was accounted for by a combination of frontal and
parietal lesions. Prefrontal cortex lesions also produce
deficits in response inhibition tasks (Grant and Berg,
1948; Malloy et al., 1993; Perret, 1974). Casey et al.
(1997) reported that ventral and dorsal prefrontal re-
gions were activated in both children and adults during
a Go/No-Go response inhibition task, although children
activated more in these regions and performed more
poorly than adults. Response inhibition is an impor-
tant component of the Stroop task, as subjects must
inhibit an inappropriate response (i.e., reading the
word) in order to correctly respond (i.e., naming the
color).
Another important cognitive process involved in
Stroop task performance is word reading and produc-
tion. Activation in prefrontal regions described in this
study also have been observed in tasks that require
vocal word reading (Bookheimer et al., 1995; Hirano et
al., 1996) as well as silent word reading (Bookheimer et
al., 1995; Hirano et al., 1996; Petersen et al., 1990).
This is an intriguing finding considering the positive
correlation between age and word reading raw score on
the behavioral Stroop task. Word reading and produc-
tion play an important role in Stroop interference res-
olution and functional development in the prefrontal
71
cortex may influence both reading skills and interfer-
ence resolution skills.
Parietal and Parieto-occipital Cortex
There was a significant linear correlation between
activation in left inferior parietal/parieto-occipital re-
gions and age. In the between-group analysis, signifi-
cant differences were confined to the adult– child and
adolescent– child comparisons. No differences in pari-
etal cortex activation, however, were evident in the
adult–adolescent comparison. This finding suggests
that Stroop task-related processes of the parietal lobe
are mostly developed by adolescence.
Activation in the parietal lobe during the Stroop task
was observed bilaterally in the inferior parietal lobule
in agreement with previous results (Carter et al.,
1995), as well as in parieto-occipital regions. Our find-
ings also are in accord with those of Bench et al. (1993)
and Peterson et al. (1999) who found Stroop task-re-
lated inferior parietal activation. These authors sug-
gest that the parietal cortex plays a role in maintaining
sustained attention during the Stroop task. In the
present study, parietal cortex activation also may re-
flect sustained attention to an important stimulus fea-
ture in the face of more salient and misleading stimu-
lus features. A number of studies have shown that
parietal regions are activated during sustained atten-
tion, attention shifts, and when attention is reflexively
drawn to prominent features of a stimulus (see
Chelazzi and Corbetta (2000) for a review). For exam-
ple, Menon et al. (1997) found both fMRI and EEG
evidence that the inferior parietal cortex plays an im-
portant role in allocation of directed attention to an
unexpected event.
In addition to playing a role in sustained attention,
parieto-occipital activity may be associated with word
reading (Brunswick et al., 1999; Buchel et al., 1998;
Shaywitz et al., 1998) and visual attention processes
(Carpenter et al., 2000). In a recent review of PET
studies, Posner et al. (1999) reported that letter strings
activate prestriate and parietal areas in the right
hemisphere, but only word-like stimuli activate these
regions in the left hemisphere. In the present study,
the presence of words in the experimental, but not the
control condition could explain the left prestriate and
parietal region activation evident in Stroop task per-
formance. In addition, previous studies suggest that
extrastriate occipital regions, which were active in the
adult– child comparison, may be involved in visual
word form analysis (Petersen et al., 1988; Pugh et al.,
1996). As cognitive development occurs and subjects
become more familiar with words and reading, it ap-
pears that they exhibit more activation in these pari-
etal areas. Reading skill may be a factor in the activa-
tion levels seen in the parietal and parieto-occipital
regions during performance of the Stroop task.
72 ADLEMAN ET AL.

FIG. 6. Only the left middle frontal gyrus and anterior cingulate cortex showed significantly greater Stroop task-related activation in the
young adults, compared to the adolescents. There were no differences in the parietal lobe. Other details as described in the legend of Fig. 2.

Anterior Cingulate Cortex Stroop task-related activation in the ACC is in agree-

The present study provides evidence for Stroop task-
related functional development in the left anterior cin-
gulate cortex. There was a significant positive linear
correlation between activation in the left ACC and age.
Greater activation in this region also was evident in
the adult–adolescent and adult– child comparisons.
These differences suggest that functional development
in the left ACC continues throughout childhood, ado-
lescence, and into young adulthood.
ment with several studies of interference resolution
(Bench et al., 1993; Bush et al., 1998; Carter et al.,
1995; Pardo et al., 1990). However, compared to Stroop
task-related activation in the lateral prefrontal cortex,
activation in the anterior cingulate cortex has been
observed less consistently. A PET study conducted by
Pardo et al. (1990) showed that the most robust acti-
vation during a Stroop task was in the ACC. Taylor et
al. (1997) observed that the ACC was active in one of
two different Stroop conditions: the Stroop compared to
 DEVELOPMENTAL fMRI OF THE STROOP TASK 73

FIG. 7. Brain areas showing significantly greater Stroop task-related activation in the adolescents, compared to the children, included
the left parietal and parieto-occipital regions such as the cuneus and posterior cingulate/precuneus. Other details as described in the legend
of Fig. 2.

neutral word contrast, but not in the Stroop compared
to false font comparison. Bench et al. (1993) found
activation in the ACC in a Stroop versus colored
crosses comparison. However, they did not find a sig-
nificant correlation between reaction times and re-
gional cerebral blood flow in the ACC and therefore,
concluded that activation in the ACC alone could not
explain Stroop interference. Activation in the ACC also
has been observed during the Go/No-Go response inhi-
bition task in adults (Menon et al., 2001) as well as in
children (Casey et al., 1997). An event-related Go/
No-Go task, however, did not yield ACC activation
(Garavan et al., 1999).
Neuroimaging studies have suggested that the ACC
may be involved in executive control processes such as
inhibition of inappropriate responses (Paus et al.,
1993), selective attention (Pardo et al., 1990), and mon-
itoring of response conflict (Barch, 1999). The ACC has
been shown to be active when subjects are instructed to
produce a less practiced response instead of a more
74 ADLEMAN ET AL.
automatic, prepotent response (Barch et al., 2000).
These executive control processes are important in the
Stroop task. In order to perform the Stroop task, sub-
jects must inhibit the inappropriate response of word
reading, pay selective attention to color, and monitor
the response conflict between the possible reponses of
color naming and word reading.
Conclusions
The results of the present study indicate that in-
creased activation occurs with age in a number of brain
regions thought to be important in performance of the
Stroop task, including the left lateral prefrontal cortex,
left parietal/parieto-occipital cortex, and left anterior
cingulate cortex. Specifically, activity in the left lateral
prefrontal cortex and the anterior cingulate increases
linearly with age, suggesting an increase in focal re-
source recruitment during maturation. This age-re-
lated increase in activation occurs in conjunction with
better behavioral performance on some measures of
the Stroop task.
The present study is the first developmental fMRI
study of the color–word Stroop test. As such, the re-
sults are preliminary and warrant further investiga-
tion. In order to minimize head movement during the
scan, we asked the subjects to silently respond to the
Stroop task presented within the scanner. Conse-
quently, a limitation of the study is that task compli-
ance and behavioral performance during the scan could
not be measured. Future studies should attempt to
record verbal responses during fMRI acquisition. More
importantly, the differentiation between verbal and
nonverbal components involved in the development of
interference processing remains to be elucidated. The
present study suggests that brain regions involved in
word reading and interference resolution processes co-
evolve with development. Future studies should at-
tempt to isolate interference processing from word
reading in a developmental paradigm.
ACKNOWLEDGMENTS
This work was supported by NIH Grants MH01142, HD31715, and
HD40761, the Packard Foundation and the Sinclair Fund.
REFERENCES
Achenbach, T. M. 1991. Manual for the Child Behavior Checklist/
4-18 and 1991 Profile. University of Vermont Department of Psy-
chiatry Press, Burlington, VT.
Barch, D. M. 1999. Images in neuroscience. Cognition: The anterior
cingulate and response conflict. Am. J. Psychiatry 156(12): 1849.
Barch, D. M., Braver, T. S., Sabb, F. W., and Noll, D. C. 2000.
Anterior cingulate and the monitoring of response conflict: Evi-
dence from an fMRI study of overt verb generation. J. Cogn.
Neurosci. 12(2): 298 –309.
Bench, C. J., Frith, C. D., Grasby, P. M., Friston, K. J., Paulesu, E.,
Frackowiak, R. S., and Dolan, R. J. 1993. Investigations of the
functional anatomy of attention using the Stroop test. Neuropsy-
chologia 31(9): 907–922.
Bettner, L. G., Jarvik, L. F., and Blum, J. E. 1971. Stroop color-word
test, non-psychotic organic brain syndrome, and chromosome loss
in aged twins. J. Gerontol. 26(4): 458 – 469.
Bonino, S., and Ciairano, S. 1997. Effetto Stroop e capacita di inibi-
zione: Una ricerca in campo evolutivo. Giornale Italiano di Psico-
logia 24(3): 587– 603.
Bookheimer, S. Y., Zeffiro, T. A., Blaxton, T., Gaillard, W. D., and
Theodore, W. H. 1995. Regional cerebral blood flow during object
naming and word reading. Hum. Brain Mapp. 3: 93–106.
Brett, M. 2000. http://www.mrc-cbu.cam.ac.uk/Imaging/mnispace.html.
Brunswick, N., McCrory, E., Price, C. J., Frith, C. D., and Frith, U.
1999. Explicit and implicit processing of words and pseudowords
by adult developmental dyslexics: A search for Wernicke’s Worts-
chatz? Brain 122(Pt. 10): 1901–1917.
Buchanan, C. M., Eccles, J. S., and Becker, J. B. 1992. Are adoles-
cents the victims of raging hormones: Evidence for activational
effects of hormones on moods and behavior at adolescence. Psychol.
Bull. 111(1): 62–107.
Buchel, C., Price, C., and Friston, K. 1998. A multimodal language
region in the ventral visual pathway. Nature 394(6690): 274 –277.
Bush, G., Whalen, P. J., Rosen, B. R., Jenike, M. A., McInerney, S. C.,
and Rauch, S. L. 1998. The counting Stroop: An interference task
specialized for functional neuroimaging—validation study with
functional MRI. Hum. Brain Mapp. 6(4): 270 –282.
Cabeza, R., and Nyberg, L. 2000. Imaging cognition II: An empirical
review of 275 PET and fMRI studies. J. Cogn. Neurosci. 12(1):
1– 47.
Carpenter, P. A., Just, M. A., and Reichle, E. D. 2000. Working
memory and executive function: Evidence from neuroimaging.
Curr. Opin. Neurobiol. 10(2): 195–199.
Carter, C. S., Mintun, M., and Cohen, J. D. 1995. Interference and
facilitation effects during selective attention: An H215O PET
study of Stroop task performance. Neuroimage 2(4): 264 –272.
Casey, B. J., Trainor, R. J., Orendi, J. L., Schubert, A. B., Nystrom,
L. E., Giedd, J. N., Castellanos, F. X., Haxby, J. V., Noll, D. C.,
Cohen, J. D., Forman, S. D., Dahl, R. E., and Rapoport, J. L. 1997.
A developmental functional MRI study of prefrontal activation
during performance of a go-no-go task. J. Cogn. Neurol. 9(6):
835– 847.
Chelazzi, L., and Corbetta, M. 2000. Cortical mechanisms of visuo-
spatial attention in the primate brain. In The New Cognitive
Neurosciences (M. S. Gazzaniga, Ed.), 2nd ed., pp. 667– 686. The
MIT Press, Cambridge, MA.
Cohen, J. D., MacWhinney, B., Flatt, M., and Provost, J. 1993.
PsyScope: A new graphic interactive environment for designing
psychology experiments. Behav. Res. Methods Instr. Comp. 25(2):
257–271.
Comalli, P. E., Jr., Wapner, S., and Werner, H. 1962. Interference
effects of Stroop color-word test in childhood, adulthood, and ag-
ing. J. Genet. Psychol. 100: 47–53.
Dash, J., and Dash, A. S. 1982. Cognitive developmental studies of
the Stroop phenomena: Cross-sectional and longitudinal data. In-
dian Psychol. 1(1): 24 –33.
Derogatis, L. R., and Savitz, K. L. 1999. The SCL-90-R, Brief Symp-
tom Inventory, and Matching Clinical Rating Scales, 2nd ed. Law-
rence Erlbaum, Mahwah, NJ.
Diamond, A., and Taylor, C. 1996. Development of an aspect of
executive control: Development of the abilities to remember what
I said and to “do as I say, not as I do.” Dev. Psychobiol. 29(4):
315–334.
Friston, K. J., Holmes, A. P., Worsley, J.-P., Poline, C. D., Frith,
C. D., and Frackowiak, R. S. J. 1995. Statistical parametric maps
 DEVELOPMENTAL fMRI OF THE STROOP TASK
in functional imaging: A general linear approach. Hum. Brain
Mapp. 2: 189 –210.
Gaillard, W. D., Hertz-Pannier, L., Mott, S. H., Barnett, A. S., LeBi-
han, D., and Theodore, W. H. 2000. Functional anatomy of cogni-
tive development: fMRI of verbal fluency in children and adults.
Neurology 54(1): 180 –185.
Garavan, H., Ross, T. J., and Stein, E. A. 1999. Right hemispheric
dominance of inhibitory control: An event-related functional MRI
study. Proc. Natl. Acad. Sci. USA 96(14): 8301– 8306.
Gerstadt, C. L., Hong, Y. J., and Diamond, A. 1994. The relationship
between cognition and action: Performance of children 3 1/2–7
years old on a Stroop-like day-night test. Cognition 53(2): 129 –
153.
Giedd, J. N., Blumenthal, J., Jeffries, N. O., Castellanos, F. X., Liu,
H., Zijdenbos, A., Paus, T., Evans, A. C., and Rapoport, J. L. 1999.
Brain development during childhood and adolescence: A longitu-
dinal MRI study [letter]. Nat. Neurosci. 2(10): 861– 863.
Glover, G. H., and Lai, S. 1998. Self-Navigated spiral fMRI: Inter-
leaved versus single-shot. Magn. Res. Med. 39(3): 361–368.
Golden, C. J. 1978. Stroop color and word test. In A Manual for
Clinical & Experimental Users. Stoelting, Wood Dale, IL.
Grant, D. A., and Berg, E. A. 1948. A behavioral analysis of degree of
reinforcement and ease of shifting to new responses in a Weigl type
card sorting problem. J. Exp. Psychol. 38: 404 – 411.
Hayes, C., and Mathias, C. 1996. Improved Brain Coil for fMRI and
High Resolution Imaging. Paper presented at the ISMRM 4th
Annual Meeting Proceedings, New York.
Hirano, S., Kojima, H., Naito, Y., and Honjo, I. 1996. Cortical speech
processing mechanisms while vocalizing visually-presented lan-
guages. Neuroreport 8(1): 363–367.
Holmes, A. P., and Friston, K. J. 1998. Generalisability, random
effects and population inference. Neuroimage 7(4): S754.
Kraemer, H. C., Yesavage, J. A., Taylor, J. L., and Kupfer, D. 2000.
How can we learn about developmental processes from cross-sec-
tional studies, or can we? Am. J. Psychiatry 157(2): 163–171.
Larrue, V., Celsis, P., Bes, A., and Marc-Vergnes, J. P. 1994. The
functional anatomy of attention in humans: Cerebral blood flow
changes induced by reading, naming, and the Stroop effect.
J. Cereb. Blood Flow Metab. 14(6): 958 –962.
MacLeod, C. M. 1991. Half a century of research on the Stroop effect:
An integrative review. Psychol. Bull. 109(2): 163–203.
Malloy, P., Bihrlr, A., and Duffy, J. 1993. The orbitomedial frontal
syndrome. Arch. Clin. Neuropsychol. 8: 185–201.
McGrew, K. S., and Woodcock, R. W. 2001. Technical Manual, Wood-
cock-Johnson III. Riverside, Itasca, IL.
Menon, V., Adleman, N. E., White, C. D., Glover, G. H., and Reiss,
A. L. 2001. Error-related brain activation during a Go/NoGo re-
sponse inhibition task. Hum. Brain Mapp. 12(3): 131–143.
Menon, V., Ford, J. M., Lim, K. O., Glover, G. H., and Pfefferbaum,
A. 1997. Combined event-related fMRI and EEG evidence for tem-
poral-parietal cortex activation during target detection. Neurore-
port 8(14): 3029 –3037.
Pardo, J. V., Pardo, P. J., Janer, K. W., and Raichle, M. E. 1990. The
anterior cingulate cortex mediates processing selection in the
Stroop attentional conflict paradigm. Proc. Natl. Acad. Sci. USA
87(1): 256 –259.
Paus, T., Petrides, M., Evans, A. C., and Meyer, E. 1993. Role of the
human anterior cingulate cortex in the control of oculomotor, man-
ual, and speech responses: A positron emission tomography study.
J. Neurophysiol. 70(2): 453– 469.
75
Perret, E. 1974. The left frontal lobe of man and the suppression of
habitual responses in verbal categorical behavior. Neuropsycholo-
gia 12: 323–330.
Petersen, S. E., Fox, P. T., Posner, M. I., Mintun, M., and Raichle,
M. E. 1988. Positron emission tomographic studies of the cortical
anatomy of single-word processing. Nature 331(6157): 585–589.
Petersen, S. E., Fox, P. T., Snyder, A. Z., and Raichle, M. E. 1990.
Activation of extrastriate and frontal cortical areas by visual
words and word-like stimuli. Science 249(4972): 1041–1044.
Peterson, B. S., Skudlarski, P., Gatenby, J. C., Zhang, H., Anderson,
A. W., and Gore, J. C. 1999. An fMRI study of Stroop word-color
interference: Evidence for cingulate subregions subserving multi-
ple distributed attentional systems. Biol. Psychiatry 45(10): 1237–
1258.
Poline, J. B., Worsley, K. J., Evans, A. C., and Friston, K. J. 1997.
Combining spatial extent and peak intensity to test for activations
in functional imaging. Neuroimage 5(2): 83–96.
Posner, M. I., Abdullaev, Y. G., McCandliss, B. D., and Sereno, S. C.
1999. Neuroanatomy, circuitry and plasticity of word reading [pub-
lished erratum appears in Neuroreport 1999 Jun 23;10(9):inside
back cover]. Neuroreport 10(3): R12–R23.
Pugh, K. R., Shaywitz, B. A., Shaywitz, S. E., Constable, R. T.,
Skudlarski, P., Fulbright, R. K., Bronen, R. A., Shankweiler, D. P.,
Katz, L., Fletcher, J. M., and Gore, J. C. 1996. Cerebral organiza-
tion of component processes in reading. Brain 119(Pt. 4): 1221–
1238.
Pujol, J., Vendrell, P., Deus, J., Junque, C., Bello, J., Marti-Vilalta,
J. L., and Capdevila, A. 2000. The effect of medial frontal and
posterior parietal demyelinating lesions on Stroop Interference.
Neuroimage, published electronically.
Rosinski, R. R. 1977. Picture-word interference is semantically
based. Child Dev. 48(2): 643– 647.
Rosinski, R. R., Golinkoff, R. M., and Kukish, K. S. 1975. Automatic
semantic processing in a picture-word interference task. Child
Dev. 46(1): 247–253.
Rubia, K., Overmeyer, S., Taylor, E., Brammer, M., Williams, S. C.,
Simmons, A., Andrew, C., and Bullmore, E. T. 2000. Functional
frontalisation with age: Mapping neurodevelopmental trajectories
with fMRI. Neurosci. Biobehav. Rev. 24(1): 13–19.
Shaywitz, S. E., Shaywitz, B. A., Pugh, K. R., Fulbright, R. K.,
Constable, R. T., Mencl, W. E., Shankweiler, D. P., Liberman,
A. M., Skudlarski, P., Fletcher, J. M., Katz, L., Marchione, K. E.,
Lacadie, C., Gatenby, C., and Gore, J. C. 1998. Functional disrup-
tion in the organization of the brain for reading in dyslexia. Proc.
Natl. Acad. Sci. USA 95(5): 2636 –2641.
Sowell, E. R., Thompson, P. M., Holmes, C. J., Jernigan, T. L., and
Toga, A. W. 1999. In vivo evidence for post-adolescent brain mat-
uration in frontal and striatal regions [letter]. Nat. Neurosci. 2(10):
859 – 861.
Taylor, S. F., Kornblum, S., Lauber, E. J., Minoshima, S., and Ko-
eppe, R. A. 1997. Isolation of specific interference processing in the
Stroop task: PET activation studies. Neuroimage 6(2): 81–92.
Thomas, K. M., King, S. W., Franzen, P. L., Welsh, T. F., Berkowitz,
A. L., Noll, D. C., Birmaher, V., and Casey, B. J. 1999. A develop-
mental functional MRI study of spatial working memory. Neuro-
image 10(3 Pt. 1): 327–338.
Vendrell, P., Junque, C., Pujol, J., Jurado, M. A., Molet, J., and
Grafman, J. 1995. The role of prefrontal regions in the Stroop task.
Neuropsychologia 33(3): 341–352.
Worsley, K. J., and Friston, K. J. 1995. Analysis of fMRI time-series
revisited—Again. Neuroimage 2(3): 173–181.